1urn:lsid:arphahub.com:pub:f9b2e808-c883-5f47-b276-6d62129e4ff4urn:lsid:zoobank.org:pub:245B00E9-BFE5-4B4F-B76E-15C30BA74C02Biodiversity Data JournalBDJ1314-28361314-2828Pensoft Publishers10.3897/BDJ.5.e964996494277urn:lsid:arphahub.com:pub:5d52f9c1-2885-556f-8df2-b28817179d67urn:lsid:zoobank.org:pub:899507E1-4FD9-44DC-9C16-782F9E738704http://tb.plazi.org/GgServer/summary/3952694AFF94A5573067E760536BFFC0Taxonomic PaperTachinidaeBiodiversity & ConservationSystematicsCosta RicaCentral America and the CaribbeanCentral AmericaAmericasNine new species of Uramya Robineau-Desvoidy (Diptera: Tachinidae) from Area de Conservación Guanacaste in northwestern Costa Rica, with a key to their identificationFlemingAJajfleming604@gmail.com1WoodD. Monty1SmithM. Alex2HallwachsWinnie3JanzenDaniel3DapkeyTanya31Agriculture Agri-Food Canada, Ottawa, CanadaAgriculture Agri-Food CanadaOttawaCanada2Department of Integrative Biology and the Biodiversity Institute of Ontario, Guelph, CanadaDepartment of Integrative Biology and the Biodiversity Institute of OntarioGuelphCanada3University of Pennsylvania, Philadelphia, United States of AmericaUniversity of PennsylvaniaPhiladelphiaUnited States of America

Corresponding author: AJ Fleming (ajfleming604@gmail.com).

Academic editor: Daniel Whitmore

2017070320175e96492206201602032017AJ Fleming, D. Monty Wood, M. Alex Smith, Winnie Hallwachs, Daniel Janzen, Tanya DapkeyThis is an open access article distributed under the terms of the Creative Commons Attribution License 4.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.AbstractBackground

We describe nine new species in the genus Uramya Robineau-Desvoidy, 1830 from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. All species were reared from an ongoing inventory of wild-caught caterpillars spanning a variety of families (Lepidoptera: Erebidae; Limacodidae; Megalopygidae; Lasiocampidae and Dalceridae). Our study provides a concise description of each new species using morphology, life history, molecular data, and photographic documentation. In addition to the new species the authors provide a redescription the previously described Uramyasibinivora Guimarães, which was also collected within ACG during this study. We also provide a redescription of the genus, and a revised key to species of Uramya occurring in Central and South America.

The following are proposed by Wood as new synonyms of Uramya: Olinda Townsend, syn. nov. and Procleonice Townsend, syn. nov. The following new combination is proposed as a result of the new synonymies: Uramyabrasiliensis Macquart, comb. nov.Procleoniceprolixa Townsend is synonymized under Uramyabrevicauda Curran, syn. nov.

Tachinidae are the second most diverse family of Diptera (Belshaw 1993, O’Hara 2008), with almost 10,000 described species classified into over 1500 genera (O’Hara 2008, O’Hara 2014). The number of named species catalogued by Guimaraes (1971) for the Neotropical Region, some 2,864 species, is larger than that of any other region. Based on what has recently been discovered in Costa Rica and what is already present in other collections, this number is undoubtedly just a small fraction of what actually exists in nature. The most speciose genera of the Costa Rican tachinid fauna seem to occur in the upper elevations and cloud forests that extend from the western slopes of the Sierra Madre Occidental in Mexico to both slopes of the Andes, from Colombia south to Bolivia. The present study describes 9 new Neotropical species of Uramya Robineau-Desvoidy, 1830 (Dexiinae: Uramyini) from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica (http://www.acguanacaste.ac.cr) and provides a key to their identification and that of their Central and South American congeners.

The last major taxonomic work on the Uramyini was by Guimarães (1980): his "Revision of the South American Uramyini (Diptera, Tachinidae)" provided a concise and complete diagnosis of the tribe and included two nominal genera: Uramya and Thelairaporia Guimarães, 1980. Robineau-Desvoidy (1830) erected the genus Uramya based on one male collected in Brasil, misidentified as a female, which he named U.producta Robineau-Desvoidy, 1830. Like other genera within the Uramyiini (e.g., Itaplectops Townsend, 1927 and Thelairaporia), species of Uramya parasitize caterpillars within the Arctiinae (Erebidae), Limacodidae, Megalopygidae, Lasiocampidae, and Dalceridae (Arnaud 1978, Guimarães 1980, Wood and Zumbado 2010).

Our descriptions of these nine new species of Uramya build on existing knowledge and are based on differences in external morphology, COI (coxI or cytochrome c oxidase I) gene sequences, and male terminalia (when necessary). As the inventory is continually growing, it should be noted that this paper should not be taken as an indication of the final total number of species of Uramya present in ACG or Costa Rica. Our descriptions are limited only to the species known and reared from ACG. This paper on Uramya is part of a larger effort to describe new species reared during the ACG inventory (Fleming et al. 2014a, Fleming et al. 2014b, Fleming et al. 2015a, Fleming et al. 2015c, Fleming et al. 2015b, Fleming et al. 2015d, Fleming et al. 2016a, Fleming et al. 2016b). This series of taxonomic papers will represent a baseline for later, detailed ecological and behavioral accounts and studies extending across ACG ecological groups, whole ecosystems, and taxonomic assemblages much larger than a genus.

Materials and methodsProject aims and rearing intensity

All reared specimens were obtained from host caterpillars collected in ACG (Janzen et al. 2009, Janzen and Hallwachs 2011, Janzen and Hallwachs 2015). ACG's 125,000+ terrestrial hectares span the provinces of Alajuela and Guanacaste, along the dry forested northwestern coast of Costa Rica and inland to the Caribbean lowland rain forest. ACG comprises several different biomes and intergrades, ranging from sea level up to 2,000 m. The tachinid rearing methods are described at http://janzen.bio.upenn.edu/caterpillars/methodology/how/parasitoid_husbandry.htm. Since its inception, this inventory has reared over 750,000 wild-caught ACG caterpillars. Any frequencies of parasitization reported here need to be considered against this background inventory. Comparative details of the parasitization ecology of these flies will be treated separately in later papers, in the context of the study of all parasitization rates of tachinids on ACG caterpillars, once the overall alpha taxonomy of ACG caterpillar-attacking tachinids is more complete than at present.

Descriptions and imaging

Species accounts presented in this paper are deliberately brief and only include basic descriptions of body morphology and coloration commonly used in the identification of Tachinidae. The descriptions are complemented with a series of color photos of every species, used to illustrate the morphological differences among them. The morphological terminology used follows Cumming and Wood (2009). All dissections and photography were carried out following the methods detailed in Fleming et al. (2014a). Measurements and examples of parts of the terminalia are illustrated in Fig. 1.

Voucher specimen management

All caterpillars reared from the ACG efforts receive a unique voucher code in the format yy–SRNP–xxxxx. Any parasitoid emerging from a caterpillar receives the same voucher code as a record of the rearing event. If and when the parasitoid is later dealt with individually it receives a second voucher code unique to it, in the format DHJPARxxxxxxx. These voucher codes assigned to both host and parasitoids may be used to obtain the individual rearing record at http://janzen.bio.upenn.edu/caterpillars/database.lasso.

To date, all DHJPARxxxxxx-coded tachinids have had one leg removed for DNA barcoding at the Biodiversity Institute of Ontario (BIO) in Guelph, ON, Canada. All successful barcodes and collateral data are first deposited in the Barcode of Life Data System (BOLD, www.boldsystems.org) (Ratnasingham and Hebert 2007), and later migrated to GenBank. Each barcoded specimen is also assigned unique accession codes from both the Barcode of Life Data System (BOLD) and GenBank respectively.

Inventoried Tachinidae were collected under Costa Rican government research permits issued to DHJ, and exported from Costa Rica to Philadelphia, en route to their final depository in the Canadian National Insect collection in Ottawa, Canada (CNC). Tachinid identifications for the inventory were done by DHJ in coordination with a) visual inspection by AJF and DMW, b) DNA barcode sequence examination by MAS and DHJ, and c) correlation with host caterpillar identifications by DHJ and WH through the inventory itself. Dates of collection cited for each ACG specimen are the dates of eclosion of the fly, not the date of capture of the caterpillar, since the fly eclosion date is much more representative of the time when that fly species is on the wing than is the time of capture of the host caterpillar. The collector listed on the label is the parataxonomist who found the caterpillar, rather than the person who retrieved the newly eclosed fly from its rearing container. The holotypes of the species newly described herein are all deposited at CNC.

Acronyms for depositories

AMNH American Museum of Natural History, New York, New York, USA

BMNH The Natural History Museum, London, United Kingdom

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada

USNM United States National Museum of Natural History, Washington, D.C., USA

Interim names of undescribed host species

Names of undescribed host species follow a standardized, interim naming system used for taxonomic units considered as distinct species and identified by DNA barcodes. The interim names are given in the format "Eois Janzen52" or "CaviriareginaDHJ01", where the "species epithet" is either composed of the name of the taxonomist who identified the species and a number or the name of a species-group followed by a code. This prevents confusion with already described species while maintaining traceability of each undescribed species within the ACG project.

DNA Barcoding

DNA barcodes (DNA sequences from a standardized 5’ region of the mitochondrial cytochrome c oxidase I (COI) gene) for all ACG inventoried specimens were obtained using DNA extracts prepared from single legs using a modified glass fibre protocol (Ivanova et al. 2006). A 658-bp region near the 5’ terminus of the COI gene was amplified from the total genomic DNA extract using standard insect primers (LepF1–LepR1 (Hebert et al. 2004)) and following established protocols (Smith et al. 2006, Smith et al. 2008).

Abdomen: narrowed and elongate, at least twice as long as wide; frequently acutely produced dorsally into a tail-like process; mid-dorsal depression on ST1+2 reaching hind margin of syntergite; T3, T4 and sometimes ST1+2 with one to three pairs of discal bristles; all tergites with marginals. In females, abdomen usually not as elongate as males and with a rounded posterior end.

Nearctic and Neotropical (not known from Chile and southern Argentina)

Ecology

According to Arnaud (1978)Uramya species parasitize lepidopteran larvae in the families Limacodidae, Arctiidae, Megalopygidae and Lasiocampidae. Current data from ACG inventoried larvae confirm this, adding records from the family Dalceridae. Uramya puparia are asymmetrical as seen in Fig. 2 (Uramyainfractasp. nov.).

Uramyaalbosetulosa can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supra-alar bristles, underside of scutellum with a tuft of white hairs near basal marginal bristle, 1 pair of median marginal bristles on ST1+2 and T3; T4 and T5 with a row of marginal bristles; underside of abdomen pale pilose and silver pollinose.

Etymology

The specific epithet is derived from the Latin adjective “ albus ”, for white, and the noun “ seta ”, for bristle, in reference to the tuft of white hairs present along the underside of the scutellum in this species.

Distribution

Costa Rica, ACG (Guanacaste and Alajuela Provinces), 340–600 m.

Ecology

Uramyaalbosetulosasp. nov. has been reared 35 times from Achariasarans (Dyar), Achariahyperoche Dognin and Achariaapicalis (Dyar) (Limacodidae), from 2,035 wild-caught mixed siblings and non-siblings found in both dry and rain forest.

Female (Fig. 6). Length: 7–10 mm. As male, except arista 1.7X as long as first flagellomere, and 3 katepisternal bristles.

Diagnosis

Uramyaconstricta can be distinguished from all other Neotropical species of Uramya by the following combination of traits: pedicel dark brown, 3 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, only 1 pair of median marginal bristles on ST1+2, T3 and T4, and by the shape of the surstylus.

Etymology

The specific epithet is derived from the Latin adjective “constrictus”, for constricted or compressed, in reference to the slightly constricted base of ST1+2 where it meets the thorax.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 1,150 m.

Ecology

Uramyaconstricta has been reared 38 times from Achariaophelians Dyar (Limacodidae) (one sibling brood of caterpillars) and once from EucleamesoamericanaDHJ04 (Lepidoptera: Limacodidae) in ACG cloud forest.

Female (Fig. 8). Length: 8–11 mm. As male, except with a slight golden tinge around the thoracic suture, one pair of median discal bristles on T3, T4 and T5, wing smoky brown translucent, only slightly infuscate around major wing veins.

Diagnosis

Uramyacontraria can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supra-alar bristles, 2 strong lateral scutellar bristles, no discal scutellar bristles, underside of scutellum with a tuft of white hairs near basal marginal bristle, abdomen flattened dorsoventrally, ST1+2 with lateral white pollinose spots on either side of mid-dorsal depression, T5 subtriangular, not strongly produced into a long, tail-like process, 1 pair of median discal bristles on T3, T4 and T5, and silver pollinosity on underside of abdomen.

Etymology

The species epithet is derived from the latin adjective " contrarius " for contrary or opposed, referring to its overall similarity but differing terminalia to U.halisidotae.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 440–520 m.

Ecology

Uramyacontraria has been reared four times from a sample of 100 wild-caught, non-sibling Caviriaregina Cramer (Erebidae, Lymantriinae) in ACG rain forest.

Uramyainfracta can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae with only a slight orange tinge and dark brown pedicel, thorax with black hairs interspersed among the bristles dorsally, and yellow-white hairs on sides and ventrally, underside of scutellum with a tuft of black hairs near basal marginal bristle, 1 pair of median marginal bristles on ST1+2, and a row of marginal bristles on T3 and T4.

Etymology

The species epithet is derived from the latin adjective " infractus " meaning broken or weakened, in reference to the broken band of silver pollinosity on T5, which appears as two pollinose spots.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 900 m.

Ecology

Uramyainfracta has been reared only once, from a total of 153 wild-caught non-sibling Natadafusca Druce (Limacodidae) caterpillars in ACG rain forest.

Uramyalativittata can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal bristle, 4–5 pairs of marginal scutellar bristles, and silver pollinosity on underside of abdomen.

Etymology

The species epithet is derived from a combination of the latin nouns " latus ", meaning side, and " vitta ", meaning band, in reference to its distinctive abdominal pattern.

Distribution

Costa Rica, ACG (Provs. Guanacaste and Alajuela), 280–700 m.

Ecology

Uramyalativittata has been reared 23 times from large caterpillars belonging to the genera Norape Walker, Megalopyge Hübner, Podalia Walker and Trosia Hübner (Megalopygidae) in a sample of 2,816 non-siblings from dry, rain and dry-rain intergrade forest.

Uramyalunula can be distinguished from all other Neotropical species of Uramya by the following combination of traits: dark brown to black antennae, 3 postsutural supra-alar bristles, 2 strong lateral scutellar bristles, weak discal scutellar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, abdomen flattened dorsoventrally, ST1+2 lacking silver pollinose spots on either side of mid-dorsal depression, T5 subtriangular, not strongly produced into a long, tail-like process, 2 pairs of median discal bristles on T3, T4 and T5, and silver pollinosity on underside of abdomen.

Etymology

The species epithet is derived from the Latin noun “lunula”, for little moon, in reference to the crescent-shaped markings present on the abdomen of this species.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 405 m.

Ecology

Uramyalunula has been reared only once, from a Podaliaorsilocha (Cramer) (Megalopygidae) caterpillar found in ACG rain forest. Podaliaorsilocha has been reared 175 times from sibling and non-sibling groups in ACG.

Uramyanitida can be distinguished from all other Neotropical species of Uramya by the following combination of traits: light-colored, orange-brown antennae, 2 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, 1 pair of median marginal bristles on ST1+2, and a row of marginal bristles on T3 and T4.

Etymology

Uramyanitida is derived from the Latin noun "nitidus", meaning bright or glossy, in reference to the glossy appearance of the abdomen under certain angles of light.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 660–1220 m.

Ecology

Uramyanitida has been reared 18 times from Parasasandrae Corrales & Epstein (Limacodidae), in a sample of 497 non-sibling, wild-caught larvae in both cloud and rain forest.

Uramyapannosa can be distinguished from all other Neotropical species of Uramya by the following combination of traits: pedicel orange, 2 postsutural supra-alar bristles, underside of scutellum with a tuft of black hairs near basal marginal bristle, overall absence of abdominal hairs, only 1 pair of median marginal bristles on ST1+2, and 2 pairs of median discal bristles on T3 and T4.

Etymology

The species epithet is derived from the Latin adjective “ pannosus ”, meaining tattered or shabby, in reference to the irregular and ragged appearance of the abdominal banding.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 900–1220 m.

Ecology

Uramyapannosa has been reared 5 times from Parasamacrodonta Hering & Hopp (Limacodidae) in both cloud and rain forest, and is the only species of tachinid to have been reared from this species of caterpillar in 166 non-sibling rearings.

Uramyapenicillata is distinguished from all other Neotropical species of Uramya by the following combination of traits: light colored pedicel, 3 postsutural supra-alar bristles, the first of which is 4X smaller than the second, underside of scutellum with a tuft of black hairs near basal marginal bristle, rows of marginal bristles on T3 and T4, and transverse bands of silver pollinosity extending across tergites to underside of abdomen.

Etymology

The species epithet is derived from the Latin noun “penicillus”, for paintbrush, in reference to the brush-like tuft of hairs present along the underside of the scutellum in this species.

Distribution

Costa Rica, ACG (Prov. Guanacaste), 1460 m

Ecology

Uramyapenicillata has been reared only once, from an Isochaetesdwagsi Corrales & Epstein (Limacodidae) caterpillar found in cloud forest. Isochaetesdwagsi has been reared 503 times from sibling and non-sibling groups in ACG.

Uramyasibinivora can be distinguished from all other Neotropical species of Uramya by the following combination of traits: conspicuous white pilosity covering the katepisternum, meron, and anepimeron, ventral surface of abdomen, dorsum of the thorax, scutellum white pilose, only one pair of median discal bristles on T3, and the shape of the surstylus.

Sides of thorax and underside of abdomen moderately pilose; legs densely covered in dark hair, making them appear brown; T3 and T4 densely covered in appressed hairs not more than 1/3 as long as discal bristles; row of discal bristles present on T5.

U.nitida sp. nov.

9

3katepisternal bristles

U.infracta sp. nov.

–

2katepisternal bristles

10

10

ST1+2 with gray pollinosity dorsolaterally; underside of scutellum with a tuft of black hairs near basal marginal bristle; ST1+2 with 2 strong median marginal bristles and a row of weaker marginal bristles, extending to lateral marginal bristles; T3 and T4 with a row of marginal bristles

U.constricta sp. nov.

–

Abdominal ST1+2 lacking gray pollinosity dorsolaterally; only one pair of median marginals on each of ST1+2 and T3, and T4 with a row of marginal bristles; underside of scutellum with a tuft of white hairs near basal marginal bristle

Abdominal ST1+2 lacking silver pollinose spots, but with a thin transverse band of silver pollinosity along posterior margin; wing veins not strongly infuscate, infuscation only visible on basal halves of R4+5 and M.

U.lunula sp. nov.

This key builds on the work accomplished by Guimarães (1980). As observed by Guimarães (1980), females show less morphological differences between species. As a result, our key is based on male morphological characters only. For female character states, see individual species descriptions.

Analysis

Fig. 19 is a Neighbor Joining tree (NJ) (Saitou and Nei 1987) for the Uramya holotypes reared and DNA-barcoded by this inventory to date. The DNA barcode sequences obtained from the ten species of ACG Uramya displayed the strong AT bias characteristic of insect mitochondrial DNA (mean percent GC content 30.33%, SE 0.1) and displayed no insertions or deletions. Within-species variation was low compared to between-species variation. All values of DNA barcode variation were calculated within BOLD and can be re-calculated in the future as more specimens or species are added to the DNA library.

Acknowledgements

We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (Janzen et al. 2009, Janzen & Hallwachs 2011) who found and reared the specimens used in this study, and the team of biodiversity managers who protect and manage the ACG forests that host these tachinids and their caterpillar hosts. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699, and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, Permian Global, and University of Pennsylvania (DHJ & WH), and in-kind support from INBio and the Museo Nacional de Costa Rica. This study has also been supported by the Government of Canada through its ongoing support of the Canadian National Collection, Genome Canada, the Biodiversity Institute of Ontario, and the Ontario Genomics Institute (2008–0GI–ICI–03) (MAS), and by a Discovery Grant from Natural Sciences and Engineering Research Council of Canada (MAS).

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Neighbor Joining tree (NJ – Saitou and Nei 1987) comparing the species of Uramya present in ACG. Tree based on the Kimura-2-parameter (Kimura 1980) made using MEGA6 (Tamura et al. 2013) for a single specimen from each of the Uramya species discussed here. Tip labels include: species name|sample accession|species parasitized|family of parasitized caterpillar, and the image of a male in lateral view. Each sample presented here is the holotype for the species described here, except U.sibinivora. This phenogram shows the low intra-specific and high inter-specific variation of CO1 barcode sequences.